Toward a revised checklist of the Western Palearctic butterflies, hyperlinked to the original descriptions at species, genus and family level (Lepidoptera, Papilionoidea)
Part III: Rationale and framework for the Pieridae.
Submitted: 12.xii.2025 | Accepted: 25.xii.2025 | Published online: 30.xii.2025.
DOI: 10.5281/zenodo.17818135

Michel Taymans¹ | Sylvain Cuvelier²
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¹ Clos du Moulin Royal 2 bte 02, B-6900 Marche-en-Famenne, Belgium. michel.taymans@hotmail.com
² Diamantstraat 4, B-8900 Ieper, Belgium. sylvain.cuvelier@telenet.be

Abstract
Taxonomic and nomenclatural issues concerning the butterfly family Pieridae are reviewed in light of the rules of the International Code of Zoological Nomenclature (ICZN), historical classifications, recent phylogenetic studies and current usage. Particular attention is given to the classification of subfamilies and tribes, where older names and synonymies have sometimes conflicted with prevailing usage, requiring rigorous evaluation under the Code.
At the tribal level, the status of Aporiini is discussed. The various interpretations concerning the species composition of certain genera are examined, and the decisions adopted for the compilation of the checklist are explained, particularly for the genera Leptidea, Gonepteryx and Colias. The status of the genus Belenois and its placement within the classification, including the resulting modifications to the checklist, are clarified. Finally, the generic structure within the tribe Anthocharidini is analysed.

Key words
Taxonomy — Checklist — Papilionoidea — Pieriidae — Dismorphiinae — Coliadinae — Pierinae — Aporiini — Anthocharidini — Leptidea — Leptidea morsei — Gonepteryx — Colias — Belenois — Euchloe — Iberochloe — Elphinstonia — Anthocharis — Euchloe pechi — Zegris meridionalis — Western Palearctic.

Introduction
Nomenclatural stability within Pieridae requires consistent application of the International Code of Zoological Nomenclature combined with careful consideration of historical usage and current taxonomic practice. Despite extensive study, the family remains affected by long-standing issues arising from historical name proliferation, changing generic and subfamilial concepts, and differing regional interpretations. Recent molecular studies have clarified several relationships but have also produced conflicting or incomplete results, particularly where taxon sampling is limited. The present revised checklist of Western Palearctic Pieridae Nomenclatural stability within Pieridae requires consistent application of the International Code of Zoological Nomenclature combined with careful consideration of historical usage and current taxonomic practice. Despite extensive study, the family remains affected by long-standing issues arising from historical name proliferation, changing generic and subfamilial concepts, and differing regional interpretations. Recent molecular studies have clarified several relationships but have also produced conflicting or incomplete results, particularly where taxon sampling is limited. The present revised checklist of West Palearctic Pieridae therefore follows a pragmatic approach, retaining well-established names where justified, applying ICZN provisions consistently, and documenting the rationale for each taxonomic decision. Unresolved cases are explicitly identified, providing a transparent and stable reference framework while indicating areas where further integrative research is required.

1. Classification of Tribes in the Subfamily Pierinae

1.1. Introduction
The group-family level classification adopted in the checklist (Taymans & Cuvelier 2025), encompassing superfamilies, families, subfamilies, tribes and subtribes, largely follows the most recent phylogenetic studies on Pieridae (Braby et al. 2006; Wahlberg et al. 2014; Ding & Zhang 2016; Wei et al., 2022; Kawahara et al., 2023; Carvalho et al. 2024), with the notable exception of the Aporiini. The rationale for this exception and the approach adopted are discussed below.

1.2. Context
Unfortunately, the International Code of Zoological Nomenclature (ICZN) provides no biological, morphological, phylogenetic or genetic criteria for the recognition of taxa at the group-family level. The Code regulates nomenclature rather than taxa themselves and therefore does not define what constitutes a family or how it should be delineated.
Prior to phylogenetic analyses, group-family ranks were assigned at the discretion of individual systematists, based on a variety of criteria. These could include the structure of the exoskeleton, wing venation, legs, palpi, genitalia or life-history traits such as eggs, larvae and pupae. By considering these characteristics, systematists aimed to provide a conceptual framework for grouping species progressively into increasingly smaller sets, ultimately resulting in genus-level classifications that comprised only a few species.
In contrast, contemporary phylogenetic studies adopt a more objective approach by linking taxonomic ranks to the historical timing of lineage divergences. For instance, in Kawahara et al. (2023), the temporal intervals used to define classification levels are: families between 100 and 80 Ma, subfamilies between 80 and 60 Ma, tribes between 60 and 40 Ma, and subtribes between 40 and 20 Ma. This method is therefore anchored in estimated divergence times derived from mathematical models and can be considered less subjectively influenced. These intervals do not necessarily correspond to geological epochs (e.g., the 60 to 40 Ma interval used to define tribes spans the mid-Palaeocene to mid-Eocene). Nevertheless, the advantage of this approach is that it bases classification on a consistent, objective criterion, whereas its drawback is the exclusion of other potentially informative characteristics.

1.3. Conclusion
Regarding the Aporiini/Aporiina, Kawahara et al. (2023) recognise the group’s monophyly but include it within the Pierini. Their supplementary phylogenetic tree indicates that the Aporiini/Aporiina branch diverged from the Pierini at an estimated 36 Ma. Thus, based on an estimated 4 Ma divergence interval (subjectively calculated), the Aporiini/Aporiina are treated as a subtribe.
The last author to consider Aporiini/Aporiina at the tribe level was Braby (2005) in his Provisional Checklist of Genera of the Pieridae. What is the rationale for following this relatively older study rather than more recent ones? First, from a technical perspective, available evidence strongly supports the monophyly of the group. Second, although it is represented by a single species in the European fauna, the group is widely distributed globally, encompassing over 400 species. Recognizing it as a tribe allows for further subdivision into subtribes; for example, Belenois (and closely related genera) could constitute a distinct subtribe, separate from Delias (and related genera) or other genus-level clusters.

1.4. References
Braby M. 2005. Provisional checklist of genera of the Pieridae (Lepidoptera: Papilionoidea). — Zootaxa 832: 1-16. https://doi.org/10.11646/zootaxa.832.1.1

Braby M., Vila R. & Pierce N. 2006. Molecular phylogeny and systematics of the Pieridae (Lepidoptera: Papilionoidea): higher classification and biogeography. — Zoological Journal of the Linnean Society 147: 239–275. https://doi.org/10.1111/J.1096-3642.2006.00218.X

Carvalho A., Owens H, St Laurent R., Earl C., Dexter K., Messcher R., Willmott K., Aduse-Poku K., Collins C., Homziak N., Hoshizaki S., Hsu Y., Kizhakke A., Kunte K., Martins D., Mega N. Morinaka S., Peggie D., Romanowski H., Sáfian S., Vila R., Wang H., Braby M., Espeland M., Breinholt J., Pierce N., Kawahara A., Lohman D. 2024. Comprehensive phylogeny of Pieridae butterflies reveals strong correlation between diversification and temperature. — iScience 27(4): (109336): 1-13. https://doi.org/10.1016/j.isci.2024.109336

Ding C. & Zhang Y. 2016. Phylogenetic relationships of Pieridae (Lepidoptera: Papilionoidea) in China based on seven gene fragments. — Entomological Science 20(1): 15-23. https://doi.org/10.1111/ens.12214

Kawahara A., Storer C., Carvalho A., Plotkin D., Condamine F., Braga M., Ellis E., St Laurent R., Li X., Barve V., Cai L., Earl C., Frandsen B., Owens H., Valencia-Montoya W., Aduse-Poku K., Toussaint E., Dexter K., Doleck T., Markee A., Messcher R., Nguyen Y., Badon J., Benítez H., Braby M., Buenavente P., Chan W., Collins S., Rabideau Childers R., Dankowicz E., Eastwood R., Fric Z., Gott R., Hall J., Hallwachs W., Hardy N., Hawkins Sipe R., Heath A., Hinolan J., Homziak N., Hsu Y., Inayoshi Y, Itliong M., Janzen D., Kitching I., Kunte K., Lamas G., Landis M., Larsen E., Larsen T., Leong J., Lukhtanov V., Maier C., Martinez J., Martins D., Maruyama K., Maunsell S., Mega N., Monastyrskii A., Morais A., Müller C., Naive M., Nielsen G., Padrón P., Peggie D., Romanowski H., Sáfián S., Saito M., Schröder S., Shirey V., Soltis D., Soltis P., Sourakov A., Talavera G., Vila R., Vlasanek P., Wang H., Warren A., Willmott K., Yago M., Jetz W., Jarzyna M., Breinholt J., Espeland M., Ries L., Guralnick R., Pierce N. & Lohman D. 2023. A global phylogeny of butterflies reveals their evolutionary history, ancestral hosts and biogeographic origins. — Nature Ecology & Evolution 7: 903-913. Article: https://doi.org/10.1038/s41559-023-02041-9 . Supplementary Materials: url.

Taymans M. & Cuvelier S. A dynamic checklist of the Western Palearctic butterflies hyperlinked to the original descriptions at species, genus and family level (Lepidoptera, Papilionoidea). — Archives of Western Palearctic Lepidoptera 2025(1): 1-70. https://doi.org/10.5281/zenodo.14733224

Wahlberg N., Rota J., Braby M., Pierce N. & Wheat C. 2014. Revised systematics and higher classification of pierid butterflies (Lepidoptera: Pieridae) based on molecular data. — Zoologica Scripta, 43, 641–650. https://doi.org/10.1111/zsc.12075.

Wei F., Huang W., Fang L., He B., Zhao Y., Zhang Y., Shu Z., Su C. & Hao J. 2023. Spatio-Temporal Evolutionary Patterns of the Pieridae Butterflies (Lepidoptera: Papilionoidea) Inferred from Mitogenomic Data. — Genes 14(1):72. Article: https://doi.org/10.3390/genes14010072. Supplementary Materials: url.

2. Classification within the Genus Leptidea Billberg, 1820

2.1. Reference classification
Since the discovery of a new species within the genus Leptidea by Real (1988), researchers (Lorković 1993; Mazel 2000–2002; …) have examined this genus in detail, first using classical methods (such as genitalia) and later based on DNA or karyological data, which led to the discovery of a second new, cryptic species (Dincă et al. 2011).
Entomologists are therefore confronted with newly recognised species that cannot be reliably identified in the field by their habitus. Leptidea reali Reissinger, [1990] and Leptidea juvernica Williams, 1946 can be distinguished from Leptidea sinapis (Linnaeus, 1758) by their genitalia (male or female), but L. reali and L. juvernica can only be reliably separated by barcoding.
The names assigned to these recently described species may still change, as they were previously confused with L. sinapis, a species to which numerous infraspecific taxa had been attributed.

2.2. What is the original status, authorship and date of L. morsei?
Two publications from 1882 compete for priority:
a) Leucophasia morsei Fenton, 1882. In: Ishikawa, Papilio 2(3): 35–36, fig 3-4. The publication date printed in the journal is March 1882.
b) Leptosia morsei Fenton, [1882]. In: Butler, Proceedings of the Zoological Society of London 1881: 855. The publication date printed in the journal is 15 November 1881, but the actual distribution date was very likely in 1882.
There is therefore considerable uncertainty regarding the actual distribution dates of these works. The approach adopted here is to follow Bozano et al. (2016), the most recent monograph treating the Dismorphiinae of the Palaearctic fauna.

2.3. Conclusion
In the checklist (Taymans & Cuvelier 2025), L. sinapis (Linnaeus, 1758), L. reali Reissinger, [1990], L. juvernica Williams, 1946, L. duponcheli (Staudinger, 1871) and L. morsei (Fenton, 1882) are adopted.

2.4. References
Bozano G., Coutsis J., Herman P., Allegrucci G., Cesaroni D. & Sbordoni V. 2016. Guide to the Butterflies of the Palearctic Region. Pieridae part 3, Subfamily Coliadinae (tribes Rhodocerini, Euremini and Catopsilia), Subfamily Dismorphiinae. — Milano: Omnes Artes (Ed.). 70 p.

Dincă V., Lukhtanov V., Talavera G. & Vila R. 2011. Unexpected layers of cryptic diversity in Wood White Leptidea butterflies. — Nature Communications 2(1): (324) 1-8. https://doi.org/10.1038/ncomms1329

Fenton M. [1882]. In: Butler A. G., On Butterflies from Japan; with which are incorporated Notes and Descriptions of new Species by Montague Fenton. — Proceedings of the Zoological Society of London 1881: 846-856. (url)

Fenton M. 1882. In: Ishikawa C., Notes on variations in some Japanese Lepidoptera. — Papilio, Organ of the New York Entomological Club 2(3): 35-37, fig. (url)

Lorković Z. 1993. Leptidea reali Reissinger, 1989 (=lorkovicii Real 1988), a new European species (Lepid., Pieridae). — Natura Croatica 2(1): 1-26. (url)

Mazel R. 2000. Le polymorphisme de deux « espèces-jumelles » Leptidea sinapis L. et L. reali Reissinger en France (Lepidoptera : Pieridae). — Linneana Belgica 17(7): 277-285. (url)

Mazel R. 2001. Le polymorphisme de deux « espèces-jumelles » Leptidea sinapis L. et L. reali Reissinger en France (Lepidoptera : Pieridae), Deuxième partie. — Linneana Belgica 18(1): 37-43. (url)

Mazel R. 2001. Leptidea sinapis L., 1758 – L. reali Reissinger, 1989, le point de la situation (Lepidoptera: Pieridae, Dismorphiinae). — Linneana Belgica 18(4): 199-202. (url)

Mazel R. & Eitschberger U. 2002. Répartition géographique de Leptidea sinapis (L., 1758) et L. reali Reissinger, 1989 au nord de l’Europe, en Russie et dans quelques pays d’Asie (Lepidoptera: Pieridae, Dismorphiinae). — Linneana Belgica 18(8): 373-376. (url)

Réal P. 1988. Lépidoptères nouveaux, principalement jurassiens. — Mémoires du Comité de liaison pour les Recherches Ecofaunistiques dans le Jura 4: 1-28. (url)

Taymans M. & Cuvelier S. A dynamic checklist of the Western Palearctic butterflies hyperlinked to the original descriptions at species, genus and family level (Lepidoptera, Papilionoidea). — Archives of Western Palearctic Lepidoptera 2025(1): 1-70. https://doi.org/10.5281/zenodo.14733224

3. Classification within the Genus Gonepteryx Leach, 1815

3.1. Introduction
The various species of this genus are relatively well known. Nevertheless, the specific status of the taxa from the Canary Islands remains uncertain, as the conclusions of several recent studies are somewhat divergent. The phylogenetic study of the genus by Brunton et al. (1998) concluded that three putative species occur in the Canary Islands: Gonepteryx cleobule (Hübner, [1824]), G. eversi Rehnelt, 1974 and G. palmae Stamm, 1963. Although Dapporto et al. (2022) recognise only a single species on these islands, G. cleobule, the figure depicting the haplotype network nonetheless reveals a significant separation among the three taxa proposed by Brunton et al. (1998). However, the barcoding data show a maximum p-distance of only 2.1% (corresponding to 11 mutations in the mitochondrial COI gene), based on a limited sample of 13 specimens from three islands. As no nuclear data are available, these results should be interpreted with caution.
In his monograph on the genus, Bozano et al. (2016) retain only two species for the Canary Islands, G. cleobule and G. palmae, and treat G. eversi as a subspecies of G. cleobule. Plates of male genitalia (by Coutsis J.) demonstrate that there is a significant difference in the structure of the male genitalia between G. cleobule and G. palmae. Are these differences sufficient to ensure reproductive isolation of these insular populations?

3.2. Conclusion
In the checklist (Taymans & Cuvelier 2025), G. cleobule and G. palmae are adopted for the Gonepteryx of the Canary Islands.

3.3. References
Bozano G., Coutsis J., Herman P., Allegrucci G., Cesaroni D. & Sbordoni V. 2016. Guide to the Butterflies of the Palearctic Region. Pieridae part 3, Subfamily Coliadinae (tribes Rhodocerini, Euremini and Catopsilia), Subfamily Dismorphiinae. — Milano: Omnes Artes (Ed.). 70 p.

Brunton C. & Hurst G. 1998. Mitochondrial DNA phylogeny of Brimstone butterflies (genus Gonepteryx) from the Canary Islands and Madeira. — Biological Journal of the Linnean Society 63(1): 69-79. https://doi.org/10.1111/j.1095-8312.1998.tb01639.x

Dapporto L., Menchetti M., Vodă R., Corbella C., Cuvelier S., Djemadi I., Gascoigne-Pees M., Hinojosa J., Lam N., Serracanta M., Talavera G., Dincă V. & Vila. R. 2022. Atlas of mitochondrial genetic diversity for Western Palearctic butterflies. — Global Ecology and Biogeography 31: 2184-2190. Article: https://doi.org/10.1111/geb.13579. Supplementary Materials: url.

Taymans M. & Cuvelier S. A dynamic checklist of the Western Palearctic butterflies hyperlinked to the original descriptions at species, genus and family level (Lepidoptera, Papilionoidea). — Archives of Western Palearctic Lepidoptera 2025(1): 1-70. https://doi.org/10.5281/zenodo.14733224

4. Classification within the Genus Colias Fabricius, 1807

4.1. Introduction
Colias represents a species‑rich genus with a broad global distribution, but its internal classification has long been debated.
Early work by Berger (1986) examined the genus in detail and proposed a subdivision into several subgenera based primarily on the morphology of tergite 8.
Verhulst’s extensive monograph (2000) took a complementary approach, relying predominantly on habitus and biological traits for species delimitation. While thorough in descriptive scope, this latter work did not incorporate a broader set of morphological or molecular characters, which may limit its ability to unravel evolutionary relationships and refine the systematics of the genus.
Significant nomenclatural efforts have since aimed to clarify the taxonomic foundation of the genus.
Grieshuber & Lamas (2007) compiled a synonymic list intended to provide a complete and authoritative account of all available names in Colias, acknowledging that many taxa remain provisionally interpreted in the absence of comprehensive evidence, and underscoring the difficulty of deciding whether phenotypically distinct populations represent full species or geographic races.
The annotated catalogue by Grieshuber, Worthy & Lamas (2012) further extended these efforts by examining type material from major museum collections and providing detailed taxonomic status for Old World taxa.
Prior to the advent of modern phylogenetic studies, few systematic treatments used criteria beyond morphology or ecology to differentiate species or subgenera.
Early phylogenetic work, often based on a restricted sampling of taxa and loci, produced results that were at times contradictory or unexpected with respect to subgroup delineation and evolutionary relationships.
This has highlighted the need for broader taxon sampling and multiple lines of evidence.
Only recently has a comprehensive phylogenetic study addressed this need: Mo et al. (2024) analysed a large dataset to assign confirmed species to well‑supported clades and to investigate diversification across the genus.
The clades identified in this study correspond only partially to the subgenera proposed by Berger (1986), indicating that traditional morphological classifications, while informative, do not always fully reflect evolutionary relationships.

4.2. Conclusion
The subdivision of Colias into subgenera will require a full systematic revision of the genus. The checklist (Taymans & Cuvelier 2025), will need to be updated regarding species arrangement on the basis of the results presented by Mo et al. (2024).

4.3. References
Berger L. 1986. Systématique du genre Colias F. Lepidoptera-Pieridae. — Lambillionea 86(7-8, suppl.): 1-68.

Grieshuber J. & Lamas G. 2007. A synonymic list of the genus Colias Fabricius, 1807 (Lepidoptera: Pieridae). — Mitteilungen der münchner entomologischen Gesellschaft 97:131-171. (url)

Grieshuber J., Worthy B. & Lamas. 2012. The genus Colias Fabricius, 1807. Jan Haugum’s annotated catalogue of the Old World Colias. — Tshikolovets Publications, Pardubice. 438 p.

Taymans M. & Cuvelier S. A dynamic checklist of the Western Palearctic butterflies hyperlinked to the original descriptions at species, genus and family level (Lepidoptera, Papilionoidea). — Archives of Western Palearctic Lepidoptera 2025(1): 1-70. https://doi.org/10.5281/zenodo.14733224

Verhulst J. 2000. Les Colias du Globe: Monograph of the Genus Colias. — Keltern: Geock & Evers (ed.). 264 p.

Mo S., Braga M., Lohman D., Nylin S., Moumou A., Wheat C., Wahlberg N., Wang M., Ma F., Zhang P. & Wang H. 2024. Rapid Evolution of Host Repertoire and Geographic Range in a Young and Diverse Genus of Montane Butterflies. — Systematic Biology 74(1): 141-157. https://doi.org/10.1093/sysbio/syae061

5. Classification of the Pieris napi (Linnaeus, 1758) complex

5.1. Introduction
The Pieris napi (Linnaeus, 1758) complex comprises several taxa for which species status remains to be demonstrated. A comprehensive monograph covering all potential species (Eitschberger 1983) recognised the following species for the western Palaearctic region: Pieris napi (Linnaeus, 1758), Pieris bryoniae (Hübner, [1806]), Pieris pseudorapae Verity, 1908 which consists of three subspecies, pseudorapae Verity from Lebanon, suffusa Sheljuzhko, 1931 from southern Russia, and balcana Lorkovic, 1968 from the Balkans, and Pieris segonzaci Le Cerf, 1923. This study relied on a broad set of criteria, including ecology, habitus, the morphology of legs and androconia, male genitalia, eggs, and other characters, to establish the species listed above. Reissinger (1990), in his checklist, adopted Eitschberger’s classification unchanged.
However, the results of hybridisation experiments (Lorkovic 1970; Hesselbarth et al. 1995) challenge this classification, as they showed that the Asia Minor and Caucasian taxa previously assigned to P. pseudorapae can freely hybridise with P. napi, whereas balcana cannot.
Dapporto et al. (2022), in The Atlas of Mitochondrial Genetic Diversity for Western Palearctic Butterflies, recognised the following species: P. balcana, P. bryoniae, P. napi, and P. segonzaci.
However, a recent genomic study posted as a bioRxiv preprint (Sala-Garcia et al. 2025) using broad phylogenomic and population genomic data across the Pieris napi group suggests patterns of phylogenetic relationships, population structure and ecological differentiation that may not fully align with purely mitochondrial or morphology-based taxonomies, and it highlights the need for further evaluation once the work has undergone formal peer review.

5.2. Conclusion
The checklist (Taymans & Cuvelier 2025), is aligned with the taxonomic conclusions of the most recent peer-reviewed study.

5.3. References
Dapporto L., Menchetti M., Vodă R., Corbella C., Cuvelier S., Djemadi I., Gascoigne-Pees M., Hinojosa J., Lam N., Serracanta M., Talavera G., Dincă V. & Vila. R. 2022. Atlas of mitochondrial genetic diversity for Western Palearctic butterflies. — Global Ecology and Biogeography 31: 2184-2190. Article: https://doi.org/10.1111/geb.13579. Supplementary Materials: url.

Eitschberger U. 1983. Systematische Untersuchungen am Pieris napi-bryoniae-Komplex (s.l.) (Lepidoptera, Pieridae). Band 1, Teil 1. — Herbipoliana, Buchreihe zur Lepidopterologie 1(1): 1-504. (url)

Eitschberger U. 1983. Systematische Untersuchungen am Pieris napi-bryoniae-Komplex (s.l.) (Lepidoptera, Pieridae). Band 1, Teil 2. — Herbipoliana, Buchreihe zur Lepidopterologie 1(2): 1-601. (url)

Hesselbarth G., van Oorschot H. & Wagener S. 1995. Die Tagfalter der Türkei: Under Berücksichtigung der Angrenzende Länder. 3 Vol. — Bocholt: Selbstverlag S. Wagener (Ed.). 2199 p.

Lorković Z. 1968[1969]. Karyologischer Beitrag zur Frage der Fortpflanzungsverhältnisse südeuropäischer Taxone von Pieris napi (L.) (Lep., Pieridae). — Bioloski Glasnik 21(1-4): 95-136.

Reissinger E. J. [1990]. Checkliste Pieridae Duponchel, 1835 (Lepidoptera) der Westpalaearktis (Europa, Nordwestafrika, Kaukasus, Kleinasien). — Atalanta 20: 149-185. (url)

Taymans M. & Cuvelier S. A dynamic checklist of the Western Palearctic butterflies hyperlinked to the original descriptions at species, genus and family level (Lepidoptera, Papilionoidea). — Archives of Western Palearctic Lepidoptera 2025(1): 1-70. https://doi.org/10.5281/zenodo.14733224

Sala-Garcia J., Vives-Ingla M., Tonzo V., Dincă V., Vila R. & Carnicer J. 2025. Phylogeography and diversification of the Pieris napi species group in the Western Palaearctic. — bioRxiv 2025.01.31.634921. (url)

6. The position of the Genus Belenois Hübner, [1819] in the classification

6.1. Introduction
The study by Kawahara A. et al. (2023) provides evidence that the genus Belenois is a member of the subtribe Aporiina. As noted in point 1 above, the status of this subtribe has been revised, and it is now recognised as the tribe Aporiini.

6.2. Conclusion
The genus Belenois Hübner, [1819], together with its subgenus Anaphaeis Hübner, [1819], and its sole species recorded in the western Palaearctic, Belenois aurota (Fabricius, 1793), should consequently be positioned in the checklist (Taymans & Cuvelier 2025), immediately after Aporia crataegi (Linnaeus, 1758).

6.3. References
Kawahara A., Storer C., Carvalho A., Plotkin D., Condamine F., Braga M., Ellis E., St Laurent R., Li X., Barve V., Cai L., Earl C., Frandsen B., Owens H., Valencia-Montoya W., Aduse-Poku K., Toussaint E., Dexter K., Doleck T., Markee A., Messcher R., Nguyen Y., Badon J., Benítez H., Braby M., Buenavente P., Chan W., Collins S., Rabideau Childers R., Dankowicz E., Eastwood R., Fric Z., Gott R., Hall J., Hallwachs W., Hardy N., Hawkins Sipe R., Heath A., Hinolan J., Homziak N., Hsu Y., Inayoshi Y, Itliong M., Janzen D., Kitching I., Kunte K., Lamas G., Landis M., Larsen E., Larsen T., Leong J., Lukhtanov V., Maier C., Martinez J., Martins D., Maruyama K., Maunsell S., Mega N., Monastyrskii A., Morais A., Müller C., Naive M., Nielsen G., Padrón P., Peggie D., Romanowski H., Sáfián S., Saito M., Schröder S., Shirey V., Soltis D., Soltis P., Sourakov A., Talavera G., Vila R., Vlasanek P., Wang H., Warren A., Willmott K., Yago M., Jetz W., Jarzyna M., Breinholt J., Espeland M., Ries L., Guralnick R., Pierce N. & Lohman D. 2023. A global phylogeny of butterflies reveals their evolutionary history, ancestral hosts and biogeographic origins. — Nature Ecology & Evolution 7: 903-913. Article: https://doi.org/10.1038/s41559-023-02041-9 . Supplementary Materials: url.

Taymans M. & Cuvelier S. A dynamic checklist of the Western Palearctic butterflies hyperlinked to the original descriptions at species, genus and family level (Lepidoptera, Papilionoidea). — Archives of Western Palearctic Lepidoptera 2025(1): 1-70. https://doi.org/10.5281/zenodo.14733224

7. Structure of the genera within the tribe Anthocharidini Scudder, 1889

7.1. Introduction
The tribe Anthocharodini constitutes a well-supported monophyletic clade, clearly distinct from the Pierini (Kawahara A.Y. et al. 2023). Nonetheless, the internal generic structure of this tribe has been the focus of several competing and markedly divergent hypotheses. Tracing back to Higgins (1975), the classification of genera, based primarily on male genitalia morphology, was as follows: Euchloe Hübner, [1819] (including tagis (Hübner, [1804])), Elphinstonia Klots, 1930, Anthocharis Boisduval, Rambur & Graslin, [1833], and Zegris Boisduval, 1836. Between 1979 and 2013, Back conducted extensive studies on the biology of numerous species within this group and published multiple papers reporting his findings, without revising the generic framework.
The studies by Back, Knebelsberger and Miller (2006, 2008), however, fundamentally reshaped the overall framework of this group. First, the genus Iberochloe Back, Knebelsberger and Miller, 2008, was established to accommodate two species, tagis Hübner, [1804], and pechi Staudinger, 1885. For clarity, a concise summary of the various proposed generic arrangements within the tribe is presented below:
Back, Knebelsberger & Miller (2008): Euchloe + ((Iberochloe + Anthocharis) + Elphinstonia)
Back, Miller & Opler (2011): Euchloe + ((Iberochloe + Elphinstonia) + Anthocharis)
Back (2020: 10): (Euchloe + (Zegris + Elphinstonia)) + (Anthocharis + Iberochloe)
Kawahara A. et al. (2023): (Anthocharis + Euchloe) + Zegris
In view of these alternative hypotheses, Marabuto et al. (2020) recommended, as a taxonomic outcome of their study, retaining Euchloe, Elphinstonia and Iberochloe as subgenera of Euchloe.
Dapporto et al. (2022) recognise only three genera, listed alphabetically as Anthocharis, Euchloe and Zegris, and treat the species assigned by Back (2020) and Marabuto et al. (2020), both at subgeneric level, to Iberochloe and Elphinstonia as members of Euchloe.

7.2. Conclusion
Views on the delimitation and hierarchical rank of genera within the tribe remain diverse and, in several cases, mutually contradictory. In the face of this ongoing instability, the checklist (Taymans & Cuvelier 2025), follows the generic and specific framework adopted by Dapporto et al. (2022), with two exceptions. The checklist (Taymans & Cuvelier 2025), includes Euchloe pechi (Staudinger, 1885), as the genetic divergence reported in that study is relatively substantial and its habitus is highly distinctive. However, this placement remains provisional, as Back (1984) demonstrated that Algerian and European populations exhibit an exceptionally high degree of reproductive compatibility, with natural copulation occurring readily and no developmental anomalies or particular susceptibilities observed in the offspring.
Back (2020) treated Zegris meridionalis (Lederer, 1853) as a valid species. In line with Dapporto et al. (2022), the present checklist (Taymans & Cuvelier 2025), considers meridionalis to be a subspecies of Zegris eupheme (Esper, [1804]).

7.3. References
Back W. 1979. Zur Biologie der europäischen und nordwestafrikanischen Populationen von Euchloe ausonia Hübner 1804 (Lep. Pieridae). — Atalanta 10(3): 225-243. (url)

Back W. 1984. Beschreibung der Präimaginalstadien von Euchloe tagis pechii Staudinger, 1885* (Lep. Pieridae). — Atalanta 15(1-2): 152-164. (url)

Back W. 1991. Taxonomische Untersuchungen innerhalb der Artengruppe um Euchloe ausonia (Hübner, 1804) (Lepidoptera, Pieridae). — Atalanta 21(3/4): 187-206. (url)

Back W. 2013. Verbreitung von Euchloe ausonia (Hübner, 1804), Euchloe daphalis (Moore, 1865) und Euchloe persica Verity, 1908 stat. nov. im Iran. — Atalanta 44: 109-117.

Back W. 2020. In: Bozano G. Guide to the Butterflies of the Palearctic Region. Pieridae part 4, Subfamily Pierinae (partim), Tribe Anthocharidini. — Milano: Omnes Artes (Ed.). 102 p.

Back W., Knebelsberger T. & Miller M.A. 2006. The phylogenetic relationships of the species and subspecies of the subgenus Elphinstonia Klots, 1930 (Lepidoptera, Pieridae). — Atalanta 37(3/4): 469-482. (url)

Back W., Knebelsberger T. & Miller M.A. 2008. Molekularbiologische Untersuchungen und Systematik der paläarktischen Arten von Euchloe Hübner, [1819] (Lepidoptera: Pieridae). — Entomologische Zeitschrift 118(4): 147-169.

Back W., Miller M. A. & Opler P. A. 2011. Genetic, Phenetic, and Distributional Relationships of Nearctic Euchloe (Pieridae, Pierinae, Anthocharidini). — The Journal of the Lepidopterists' Society 65(1): 1-14. https://doi.org/10.18473/lepi.v65i1.a1

Dapporto L., Menchetti M., Vodă R., Corbella C., Cuvelier S., Djemadi I., Gascoigne-Pees M., Hinojosa J., Lam N., Serracanta M., Talavera G., Dincă V. & Vila. R. 2022. Atlas of mitochondrial genetic diversity for Western Palearctic butterflies. — Global Ecology and Biogeography 31: 2184-2190. Article: https://doi.org/10.1111/geb.13579. Supplementary Materials: url.

Higgins L. 1975. The Classification of European Butterflies. — London: Collins (Ed.). 320 p.

Kawahara A., Storer C., Carvalho A., Plotkin D., Condamine F., Braga M., Ellis E., St Laurent R., Li X., Barve V., Cai L., Earl C., Frandsen B., Owens H., Valencia-Montoya W., Aduse-Poku K., Toussaint E., Dexter K., Doleck T., Markee A., Messcher R., Nguyen Y., Badon J., Benítez H., Braby M., Buenavente P., Chan W., Collins S., Rabideau Childers R., Dankowicz E., Eastwood R., Fric Z., Gott R., Hall J., Hallwachs W., Hardy N., Hawkins Sipe R., Heath A., Hinolan J., Homziak N., Hsu Y., Inayoshi Y, Itliong M., Janzen D., Kitching I., Kunte K., Lamas G., Landis M., Larsen E., Larsen T., Leong J., Lukhtanov V., Maier C., Martinez J., Martins D., Maruyama K., Maunsell S., Mega N., Monastyrskii A., Morais A., Müller C., Naive M., Nielsen G., Padrón P., Peggie D., Romanowski H., Sáfián S., Saito M., Schröder S., Shirey V., Soltis D., Soltis P., Sourakov A., Talavera G., Vila R., Vlasanek P., Wang H., Warren A., Willmott K., Yago M., Jetz W., Jarzyna M., Breinholt J., Espeland M., Ries L., Guralnick R., Pierce N. & Lohman D. 2023. A global phylogeny of butterflies reveals their evolutionary history, ancestral hosts and biogeographic origins. — Nature Ecology & Evolution 7: 903-913. Article: https://doi.org/10.1038/s41559-023-02041-9 . Supplementary Materials: url.

Marabuto E., Pina-Martins F., Rebelo M. & Paulo O. 2020. Ancient divergence, a crisis of salt and another of ice shaped the evolution of the west Mediterranean butterfly Euchloe tagis. — Biological Journal of the Linnean Society 131(3): 487–504. https://doi.org/10.1093/biolinnean/blaa129

Taymans M. & Cuvelier S. A dynamic checklist of the Western Palearctic butterflies hyperlinked to the original descriptions at species, genus and family level (Lepidoptera, Papilionoidea). — Archives of Western Palearctic Lepidoptera 2025(1): 1-70. https://doi.org/10.5281/zenodo.14733224

Author contribution
Michel Taymans: conceptualisation, analysis, visualisation, writing - original draft, writing – review and editing.
Sylvain Cuvelier: analysis, validation, visualisation, writing – review and editing.

Acknowledgements
We are sincerely grateful to Theo Garrevoet for his careful and thorough review of the final draft, and we wish to acknowledge Bénédicte Jonckers for her invaluable administrative support as production editor.